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adult females adult males aequatorialis Alouattinae apparently approached arboreal associated animals Azara Barro Colorado Island become behave brachiating capuchin monkeys capuchins Caraya census cent clan males close Coiba Island complemental males consorts coordination copulation crossing defensive described eaten fallen young feeding feet females and young frequently fruit function Gatun Lake groups of howlers howler clans Howler females howler groups howler population howling monkeys individuals juvenile juvenile animals leaves limb located locomotion lodge tree months mother move observed occur ocelots oestrous female oestrous period palliata Panama Paraguay period of coverage period of intromission PLATE platyrrhine play play-fighting positions postures primary forest primary sexual activity primates Puerto Armuelles react reactions Rengger responses rhythmic tongue movements roars seconds sexual behavior Shannon situations social relations socionomic specific spider monkeys stimulation subgrouping tail tendency territorial range tion vocal patterns vocalizations yards young animals young howlers